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(Arecaceae): Évolution Du Système Sexuel Et Du Nombre D'étamines

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(Arecaceae): Évolution Du Système Sexuel Et Du Nombre D'étamines Etude de l’appareil reproducteur des palmiers (Arecaceae) : évolution du système sexuel et du nombre d’étamines Elodie Alapetite To cite this version: Elodie Alapetite. Etude de l’appareil reproducteur des palmiers (Arecaceae) : évolution du système sexuel et du nombre d’étamines. Sciences agricoles. Université Paris Sud - Paris XI, 2013. Français. NNT : 2013PA112063. tel-01017166 HAL Id: tel-01017166 https://tel.archives-ouvertes.fr/tel-01017166 Submitted on 2 Jul 2014 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. UNIVERSITE PARIS-SUD ÉCOLE DOCTORALE : Sciences du Végétal (ED 45) Laboratoire d'Ecologie, Systématique et E,olution (ESE) DISCIPLINE : -iologie THÈSE DE DOCTORAT SUR TRAVAUX soutenue le ./05/10 2 par Elodie ALAPETITE ETUDE DE L'APPAREIL REPRODUCTEUR DES PAL4IERS (ARECACEAE) : EVOLUTION DU S5STE4E SE6UEL ET DU NO4-RE D'ETA4INES Directeur de thèse : Sophie NADOT Professeur (Uni,ersité Paris-Sud Orsay) Com osition du jury : Rapporteurs : 9ean-5,es DU-UISSON Professeur (Uni,ersité Pierre et 4arie Curie : Paris VI) Porter P. LOWR5 Professeur (4issouri -otanical Garden USA et 4uséum National d'Histoire Naturelle Paris) Examinateurs : Anders S. -ARFOD Professeur (Aarhus Uni,ersity Danemark) Isabelle DA9OA Professeur (Uni,ersité Paris Diderot : Paris VII) 4ichel DRON Professeur (Uni,ersité Paris-Sud Orsay) 3 4 Résumé Les palmiers constituent une famille emblématique de monocotylédones, comprenant 183 genres et environ 2500 espèces distribuées sur tous les continents dans les zones tropicales et subtropicales. Leurs feuilles et leurs stipes, très caractéristiques, les rendent facilement reconnaissables dans la nature. En revanche leurs fleurs passent souvent inaperçues. Elles sont généralement petites (quelques centimètres), trimères, unisexuées, peu colorées (blanches ou vertes) et regroupées sur de grandes inflorescences. Cependant les palmiers présentent une diversité importante au niveau du système sexuel et du nombre d’étamines. Les trois systèmes sexuels principaux des angiospermes : hermaphrodisme, monoécie et dioécie, sont présents chez les palmiers. Le nombre d’étamines varie entre quelques unités (oligandrie) et des dizaines, voire centaines, d’unités (polyandrie) chez certains genres. Nous avons étudié l’évolution du système sexuel et du nombre d’étamines à l’échelle de la famille. Nous avons pour cela utilisé une phylogénie comprenant tous les genres de palmiers, bien résolue, datée et qui a été publiée récemment. Notre étude a montré que l’ancêtre commun à tous les palmiers était probablement monoïque et possédait des fleurs oligandres à 6 étamines. A partir de ces états ancestraux, plusieurs transitions ont eu lieu : vers l’hermaphrodisme et la monoécie d’une part, et vers la polyandrie d’autre part. Dans l’objectif d’initier une recherche sur une éventuelle explication fonctionnelle de l’augmentation du nombre d’étamines, nous avons comparé celui-ci à la production de pollen, en étudiant la quantité totale de pollen produite par les fleurs de 82 espèces. Notre étude a montré que, chez deux sous-familles, les fleurs ont tendance à produire plus de pollen quand le nombre d’étamines est plus élevé. Nous avons également réalisé la phylogénie moléculaire d’une sous-tribu (les Ptychospermatinae) dans laquelle la variation du nombre d’étamines est exceptionnelle. De futures études sur la génétique, le développement, l’écologie et la biologie de la pollinisation sont nécessaires. Mots clés : Arecaceae, évolution, morphologie de la fleur, nombre d’étamines, palmiers, production de pollen, phylogénie moléculaire, système sexuel Reproductive structures in palms (Arecaceae): evolution of sexual system and stamen number Abstract Palms (Arecaceae) are an emblematic family of monocots of 183 genera and around 2500 species distributed on all continents, throughout tropical and subtropical areas. Their characteristic leaves and stems make palms immediately recognizable in the field. The inconspicuous palm flowers are usually considered as rather dull. They are usually small (a few centimetres), trimerous, often unisexual, colourless (white or greenish) and grouped into huge inflorescences. However palms exhibit a large diversity in sexual system and in stamen number, diversity that is still poorly understood. The three main sexual systems of angiosperm, hermaphroditism, dioecy and monoecy are present in palms. Stamen number ranges between a few units (oligandry) to several dozens and even several hundreds of units (polyandry) in some genera. We studied the evolution of sexual system and stamen number at the family level. We used as historical framework a well-supported and dated phylogeny, published recently. Our study showed that the putative ancestor of palms was monoecious and 5 bore oligandrous flowers with 6 stamens. From these ancestral states, several transitions occurred: towards hermaphroditism and dioecy and towards polyandry respectively. In order to initiate a research on a possible functional significance of increase in stamen number, we investigated the relationship between stamen number and pollen production, by extracting the total pollen content from flowers of 82 species. Our study showed a tendency towards higher pollen production when the number of stamen increases in two subfamilies. We also produced molecular phylogeny of a subtribe (Ptychospermatinae) in which the range of variation in stamen number is exceptional. Further investigations into genetic, developmental, ecology and pollination biology are needed. Keywords: Arecaceae, palms, sexual system, stamen number, evolution, flower morphology, pollen production, molecular phylogeny Laboratoire : UMR 8079 - Ecologie, Systématique et Evolution Université Paris-Sud, Bât. 360 91405 Orsay cedex 6 Remerciements Je tiens à remercier tout particulièrement Sophie NADOT, ma directrice de thèse, pour ses précieux conseils et encouragements ainsi que pour la généreuse confiance qu’elle m’a accordée. Merci de m’avoir permis de travailler sur les palmiers, un groupe de végétaux particulièrement intriguant et fascinant, qui a affermit mon gout pour la botanique. Je remercie Sophie également de m’avoir fait partager son expérience de l’enseignement et de m’avoir confié des TD et des TP si intéressants et enrichissants. Je remercie tous les membres de l’équipe DEV : permanents Béatrice ALBERT, Hervé SAUQUET et non permanents Boris DOMENECH, Julien MASSONI, Charlotte PRIEU, Elisabeth REYES, Franck SIMONNET pour les discussions et la belle synergie scientifique qui règne dans notre équipe. Je remercie également ceux qui sont partis vivre de nouvelles aventures, Julie SANNIER et Olivier CHAUVEAU. Ils ont beaucoup aidé la novice que j’étais en biologie moléculaire. Je remercie Christian RAQUIN et Yves LOUBLIER pour leur soutien et leur precieux conseils dans mes expériences sur le pollen. Je remercie les spécialistes des palmiers qui m’ont conseillé et accompagné durant toute cette thèse : Anders BARFOD de l’Université d’Aarhus au Danemak, Bill BAKER des Jardins Botaniques Royaux de Kew, Jean-Christophe PINTAUD, Bertha LUDENA, James TREGEAR de l’IRD de Montpellier. Je tiens à remercier chaleureusement Lauren GARDINER et Laszlo CSIBA qui m’ont aidé pour les prélèvements de matériel à Kew. Je remercie les stagiaires qui ont partagé mon travail et ont j’espère pris goût à la botanique et à la recherche, Nicolas MAINGRET, Aurélie NALIN, Vanessa VILARD, Gérard IMANI, Stéphanie VALLEE. Ils m’ont tous été d’une aide précieuse dans mes recherches. Je remercie tous les membres du labo ESE pour leur accueil et leur soutien, avec une pensée spéciale pour celles et ceux qui ont partagé leur bureau et leurs pauses thé avec moi. Je tiens finalement à remercier les membres de mon jury d’avoir accepté d’évaluer mon travail et je m’excuse pour tous ceux que j’ai sans doute oubliés. 7 8 Table des matières Introduction ..............................................................................................................................13 Partie I : L’évolution de la morphologie de l’inflorescence et de la fleur ...........................15 Les plantes et l’évolution .................................................................................................15 L’évolution des traits morphologiques.............................................................................15 Les Angiospermes............................................................................................................17 La fleur .............................................................................................................................21 Le contrôle génétique de la mise en place des organes floraux .......................................26 La morphologie de la fleur et la pollinisation ..................................................................27 Les inflorescences ............................................................................................................29 Partie II : La famille des palmiers ........................................................................................31
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